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The Cape fur seal (Arctocephalus pusillus pusillus) is one of the most colonial mammals, with colonies of up to hundreds of thousands of individuals during the breeding season. During the lactation period, mothers and pups are regularly separated as females undertake multi-day foraging trips at sea. Mothers and pups use a mutual vocal recognition system to reunite after separation. Such communication is highly constrained by both high background noise and risk of individual confusion owing to the density of seals. This study aimed to experimentally assess the acoustic features relevant for mother-pup vocal identification and the propagation properties of their calls. Playback experiments revealed that mother and pup individual vocal signatures rely on both temporal and frequency parameters: amplitude and frequency modulations, timbre and fundamental frequency (f0). This is more parameters than in any colonial species studied so far. The combinational use of acoustic features reinforces the concept that both environmental and social constraints may have acted as selective pressures on the individual vocal recognition systems. Theoretical propagation distances of mother and pup vocalisations were estimated to be below the range of distances at which mother-pup reunions can occur. This suggests that Cape fur seals may have strong abilities to extract vocal signals from the background noise, as previously demonstrated in the highly colonial king penguin. Investigating the transmission of information throughout the propagation of the signal as well as the ability of the receiving individual to decipher vocal signatures is crucial to understanding vocal recognition systems in the wild., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Immune defenses are crucial for survival but costly to develop and maintain. Increased immune investment is therefore hypothesized to trade-off with other life-history traits. Here, we examined innate and adaptive immune responses to environmental heterogeneity in wild Antarctic fur seals. In a fully crossed, repeated measures design, we sampled 100 pups and their mothers from colonies of contrasting density during seasons of contrasting food availability. Biometric and cortisol data as well as blood for the analysis of 13 immune and oxidative status markers were collected at two key life-history stages. We show that immune responses of pups are more responsive than adults to variation in food availability, but not population density, and are modulated by cortisol and condition. Immune investment is associated with different oxidative status markers in pups and mothers. Our results suggest that early life stages show greater sensitivity to extrinsic and intrinsic effectors, and that immunity may be a strong target for natural selection even in low-pathogen environments such as Antarctica., (© 2024. The Author(s).)

Inbreeding depression, the loss of offspring fitness due to consanguineous mating, is generally detrimental for individual performance and population viability. We investigated inbreeding effects in a declining population of Antarctic fur seals (Arctocephalus gazella) at Bird Island, South Georgia. Here, localised warming has reduced the availability of the seal's staple diet, Antarctic krill, leading to a temporal increase in the strength of selection against inbred offspring, which are increasingly failing to recruit into the adult breeding population. However, it remains unclear whether selection operates before or after nutritional independence at weaning. We therefore used microsatellite data from 885 pups and their mothers, and SNP array data from 98 mother-offspring pairs, to quantify the effects of individual and maternal inbreeding on three important neonatal fitness traits: birth mass, survival and growth. We did not find any clear or consistent effects of offspring or maternal inbreeding on any of these traits. This suggests that selection filters inbred individuals out of the population as juveniles during the time window between weaning and recruitment. Our study brings into focus a poorly understood life-history stage and emphasises the importance of understanding the ecology and threats facing juvenile pinnipeds., (© 2024. The Author(s).)

While the majority of studies have concluded that sleep deprivation causes detrimental effects on various cognitive processes, some studies reported conflicting results. We examined the effects of a 108-h total sleep deprivation (TSD) on working memory in the northern fur seal, an animal with unusual sleep phenomenology and long-range annual migrations. The performance of fur seals was evaluated in a two-choice visual delayed matching to sample (DMTS) task, which is commonly used to evaluate working memory. In baseline conditions, the performance of fur seals in a DMTS task based on the percentage of errors was somewhat comparable with that in nonhuman primates at similar delays. We have determined that a 108-h TSD did not affect fur seals' performance in a visual DMTS task as measured by overall percentage of errors and response latencies. On the contrary, all fur seals improved task performance over the study, including the baseline, TSD and recovery conditions. In addition, TSD did not change the direction and strength of the pattern of behavioral lateralization in fur seals. We conclude that a 108-h TSD did not interfere with working memory in a DMTS test in northern fur seals., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The ability of predators to adopt hunting tactics that minimise escape reactions from prey is crucial for efficient foraging, and depends on detection capabilities and locomotor performance of both predators and prey. Here, we investigated the efficiency of a small pinniped, the Antarctic fur seal (Arctocephalus gazella) at exploiting their small prey by describing for the first time their fine-scale predator-prey interactions. We compared these with those from another diving predator, the southern elephant seal (Mirounga leonina) that forage on the same prey type. We used data recorded by a newly developed sonar tag that combines active acoustics with ultrahigh-resolution movement sensors to study simultaneously the fine-scale behaviour of both Antarctic fur seals and prey during predator-prey interactions in more than 1200 prey capture events for eight female Antarctic fur seals. Our results showed that Antarctic fur seals and their prey detect each other at the same time, i.e. 1-2 s before the strike, forcing Antarctic fur seals to display reactive fast-moving chases to capture their prey. In contrast, southern elephant seals detect their prey up to 10 s before the strike, allowing them to approach their prey stealthily without triggering an escape reaction. The active hunting tactics used by Antarctic fur seals is probably very energy consuming compared with the stalking tactics used by southern elephant seals but might be compensated for by the consumption of faster-moving larger prey. We suggest that differences in manoeuvrability, locomotor performance and detection capacities and in pace of life between Antarctic fur seals and southern elephant seals might explain these differences in hunting styles., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Ecological theory predicts niche partitioning between high-level predators living in sympatry as a mechanism to minimise the selective pressure of competition. Accordingly, male Australian fur seals Arctocephalus pusillus doriferus and New Zealand fur seals A. forsteri that live in sympatry should exhibit partitioning in their broad niches (in habitat and trophic dimensions) in order to coexist. However, at the northern end of their distributions in Australia, both are recolonising their historic range after a long absence due to over-exploitation, and their small population sizes suggest competition should be weak and may allow overlap in niche space. We found some niche overlap, yet clear partitioning in diet trophic level (δ 15 N values from vibrissae), spatial niche space (horizontal and vertical telemetry data) and circadian activity patterns (timing of dives) between males of each species, suggesting competition may remain an active driver of niche partitioning amongst individuals even in small, peripheral populations. Consistent with individual specialisation theory, broad niches of populations were associated with high levels of individual specialisation for both species, despite putative low competition. Specialists in isotopic space were not necessarily specialists in spatial niche space, further emphasising their diverse individual strategies for niche partitioning. Males of each species displayed distinct foraging modes, with Australian fur seals primarily benthic and New Zealand fur seals primarily epipelagic, though unexpectedly high individual specialisation for New Zealand fur seals might suggest marginal populations provide exceptions to the pattern generally observed amongst other fur seals., (© 2024. Crown.)

Marine central-place foragers are increasingly faced with altered prey landscapes, necessitating predictions of the impact of such changes on behavior, reproductive success, and population dynamics. We used state-dependent behavioral life history theory implemented via Stochastic Dynamic Programming (SDP) to explore the influence of changes in prey distribution and energy gain from foraging on the behavior and reproductive success of a central place forager during lactation. Our work is motivated by northern fur seals (Callorhinus ursinus) because of the ongoing population decline of the Eastern Pacific stock and projected declines in biomass of walleye pollock (Gadus chalcogrammus), a key fur seal prey species in the eastern Bering Sea. We also explored how changes in female and pup metabolic rates, body size, and lactation duration affected model output to provide insight into traits that might experience selective pressure in response to reductions in prey availability. Simulated females adopted a central-place foraging strategy after an initial extended period spent on land (4.7-8.3 days). Trip durations increased as the high energy prey patch moved farther from land or when the energy gain from foraging decreased. Increases in trip duration adversely affected pup growth rates and wean mass despite attempts to compensate by increasing land durations. Metabolic rate changes had the largest impacts on pup wean mass, with reductions in a pup's metabolic rate allowing females to successfully forage at distances of 600+ km from land for up to 15+ days. Our results indicate that without physiological adaptations, a rookery is unlikely to be viable if the primary foraging grounds are 400 km or farther from the rookery. To achieve pup growth rates characteristic of a population experiencing rapid growth, model results indicate the primary foraging grounds need to be <150 km from the rookery., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022. Published by Elsevier Ltd.)

Determining trophic habits of predator communities is essential to measure interspecific interactions and response to environmental fluctuations. South American fur seals, Arctocephalus australis (SAFS) and sea lions Otaria byronia (SASL), coexist along the coasts of Peru. Recently, ocean warming events (2014-2017) that can decrease and impoverish prey biomass have occurred in the Peruvian Humboldt Current System. In this context, our aim was to assess the effect of warming events on long-term inter- and intra-specific niche segregation. We collected whisker from SAFS (55 females and 21 males) and SASL (14 females and 22 males) in Punta San Juan, Peru. We used δ13C and δ15N values serially archived in otariid whiskers to construct a monthly time series for 2005-2019. From the same period we used sea level anomaly records to determine shifts in the predominant oceanographic conditions using a change point analysis. Ellipse areas (SIBER) estimated niche width of species-sex groups and their overlap. We detected a shift in the environmental conditions marking two distinct periods (P1: January 2005-October 2013; P2: November 2013-December 2019). Reduction in δ15N in all groups during P2 suggests impoverished baseline values with bottom-up effects, a shift towards consuming lower trophic level prey, or both. Reduced overlap between all groups in P2 lends support of a more redundant assemblage during the colder P1 to a more trophically segregated assemblage during warmer P2. SASL females show the largest variation in response to the warming scenario (P2), reducing both ellipse area and δ15N mean values. Plasticity to adapt to changing environments and feeding on a more available food source without fishing pressure can be more advantageous for female SASL, albeit temporary trophic bottom-up effects. This helps explain larger population size of SASL in Peru, in contrast to the smaller and declining SAFS population., Competing Interests: The authors have declared that no competing interests exist.

The success of maternal foraging strategies during the rearing period can greatly impact the physiology and survival of dependent offspring. Surprisingly though, little is known on the fitness consequences of foraging strategies during the foetal period. In this study, we characterized variation in maternal foraging strategy throughout pregnancy in a marine top predator (South American fur seal, Arctocephalus australis), and asked if these shifts predicted neonatal health and postnatal survival. We found that during early pregnancy all pregnant females belonged to a single, homogenized foraging niche without evident clusters. Intriguingly though, during late pregnancy, individual fur seal mothers diverged into two distinct foraging niches characterized by a benthic-nearshore and a pelagic-offshore strategy. Females that shifted towards the benthic-nearshore strategy gave birth to pups with greater body mass, higher plasmatic levels of glucose and lower levels of blood urea nitrogen. The pups born to these benthic females were eight times more likely to survive compared to females using the pelagic-offshore foraging strategy during late pregnancy. These survival effects were mediated primarily by the impact of foraging strategies on neonatal glucose independent of protein metabolic profile and body mass. Benthic-nearshore foraging strategies during late pregnancy potentially allow for the greater maternal transfer of glucose to the foetus, leading to higher chances of neonatal survival. These results call for a deeper understanding of the balance between resource acquisition and allocation provided by distinct foraging polymorphisms during critical life-history periods, and how this trade-off may be adaptive under certain environmental conditions., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Management of gases during diving is not well understood across marine mammal species. Prior to diving, phocid (true) seals generally exhale, a behaviour thought to assist with the prevention of decompression sickness. Otariid seals (fur seals and sea lions) have a greater reliance on their lung oxygen stores, and inhale prior to diving. One otariid, the Antarctic fur seal ( Arctocephalus gazella ), then exhales during the final 50-85% of the return to the surface, which may prevent another gas management issue: shallow-water blackout. Here, we compare data collected from animal-attached tags (video cameras, hydrophones and conductivity sensors) deployed on a suite of otariid seal species to examine the ubiquity of ascent exhalations for this group. We find evidence for ascent exhalations across four fur seal species, but that such exhalations are absent for three sea lion species. Fur seals and sea lions are no longer genetically separated into distinct subfamilies, but are morphologically distinguished by the thick underfur layer of fur seals. Together with their smaller size and energetic dives, we suggest their air-filled fur might underlie the need to perform these exhalations, although whether to reduce buoyancy and ascent speed, for the avoidance of shallow-water blackout or to prevent other cardiovascular management issues in their diving remains unclear. This article is part of the theme issue 'Measuring physiology in free-living animals (Part I)'.

Tracking studies of juveniles are rare compared to those of adults, and consequently little is known about the influence of intrinsic and extrinsic factors on activity during this critical life stage. We used hourly GPS data, collected from 66 Antarctic fur seal pups from birth until moulting, to investigate the explanatory power of multiple individual-based and environmental variables on activity levels. Pups were sampled from two nearby breeding colonies of contrasting density during two subsequent years, and a two-state hidden Markov model was used to identify modalities in their movement behaviour, specifically 'active' and 'inactive' states. We found that movement was typified by central place exploration, with active movement away from and subsequent return to a location of inactivity. The probability of such directed exploration was unaffected by several factors known to influence marine mammal movement including sex, body condition, and temperature. Compared to pups born at the high-density colony, pups at low-density were more active, increased their activity with age, and transitioned earlier into the tussock grass, which offers protection from predators and extreme weather. Our study illustrates the importance of extrinsic factors, such as colony of birth, to early-life activity patterns and highlights the adaptive potential of movement., (© 2021. The Author(s).)

The diet of Antarctic fur seals (Arctocephalus gazella) at South Georgia is dominated by Antarctic krill (Euphausia superba). During the breeding season, foraging trips by lactating female fur seals are constrained by their need to return to land to provision their pups. Post-breeding, seals disperse in order to feed and recover condition; estimates indicate c.70% of females remain near to South Georgia, whilst others head west towards the Patagonian Shelf or south to the ice-edge. The krill fishery at South Georgia operates only during the winter, providing the potential for fur seal: fishery interaction during these months. Here we use available winter (May to September) tracking data from Platform Terminal Transmitter (PTT) tags deployed on female fur seals at Bird Island, South Georgia. We develop habitat models describing their distribution during the winters of 1999 and 2003 with the aim of visualising and quantifying the degree of spatial overlap between female fur seals and krill harvesting in South Georgia waters. We show that spatial distribution of fur seals around South Georgia is extensive, and that the krill fishery overlaps with small, highly localised areas of available fur seal habitat. From these findings we discuss the implications for management, and future work., Competing Interests: The authors have declared that no competing interests exist.

The skin of the South American fur seal (Arctocephalus australis) is important for animal thermoregulation in both terrestrial and aquatic environments. Skin tissue samples were collected from A. australis for microscopic analysis and were related to anatomical references. The aim of this study was to describe the skin morphology, as well as to suggest the major anatomical regions and skin components involved in the thermoregulation of this species. Using light microscopy, the skin of six animals was examined based on histological, morphometric, and immunohistochemical criteria. Hair follicle morphology was examined using scanning electron microscopy. The skin was classified as either thick or thin based on its epidermal thickness. The thin epidermis regions had more abundant hair follicles, as well as high pigmentation, whereas the thick epidermis regions had very pigmented epidermal layers. Pigmentation of hair and skin is fundamental for protection against ultraviolet rays; moreover, hair is important in preventing abrasion, and provides an insulating layer against the external environment, which can be much colder than body temperature. Furthermore, the dermis is well vascularized, especially the superficial dermis. All regions of the skin have adaptations for maintaining the animal's condition in both terrestrial and aquatic environments. Among the studied regions, the interdigital region from hindflipper showed important morphological characteristics related to thermoregulation, such as having an epidermis of intermediate thickness, a dermis with a small number of hairs, a large amount of blood vessels, and sweat glands with large lumens, indicating that heat exchange in this region may be faster., (© 2020 American Association for Anatomy.)

We present estimates of the seasonal and spatial occupation by pinnipeds of the Wildlife Refuge of Ilha dos Lobos (WRIL), based on aerial photographic censuses. Twenty aerial photographic censuses were analysed between July 2010 and November 2018. To assess monthly differences in the numbers of pinnipeds in the WRIL we used a Generalized Linear Mixed Model. Spatial analysis was carried out using Kernel density analysis of the pinnipeds on a grid plotted along the WRIL. Subadult male South American sea lions (Otaria flavescens) were the most abundant pinniped in the WRIL. Potential females of this species were also recorded during half of the census. The maximum number of pinnipeds observed in the WRIL was 304 in September 2018, including an unexpected individual southern elephant seal (Mirounga leonina), and a high number of South American fur seal yearlings (Arctocephalus australis). However, there was no statistically significant difference in counts between months. In all months analysed, pinnipeds were most often found concentrated in the northern portion of the island, with the highest abundances reported in September. This study confirms the importance of the WRIL as a haulout site for pinnipeds in Brazil, recommends that land research and recreational activities occur in months when no pinnipeds are present, and encourages a regulated marine mammal-based tourism during winter and spring months.

Understanding the factors which influence foraging behaviour and success in marine mammals is crucial to predicting how their populations may respond to environmental change. The Australian fur seal (Arctocephalus pusillus doriferus, AUFS) is a predominantly benthic forager on the shallow continental shelf of Bass Strait, and represents the greatest biomass of marine predators in south-eastern Australia. The south-east Australian region is experiencing rapid oceanic warming, predicted to lead to substantial alterations in prey diversity, distribution and abundance. In the present study, foraging effort and indices of foraging success and efficiency were investigated in 138 adult female AUFS (970 foraging trips) during the winters of 1998-2019. Large scale climate conditions had a strong influence on foraging effort, foraging success and efficiency. Foraging effort and foraging success were also strongly influenced by winter chlorophyll-a concentrations and sea-surface height anomalies in Bass Strait. The results suggest increasing foraging effort and decreasing foraging success and efficiency under anticipated environmental conditions, which may have population-level impacts.

Predators impact preyscapes (3-D distribution of forage species) by consuming prey according to their abilities or by altering prey behavior as they avoid being consumed. We elucidate prey (Antarctic silverfish[Pleuragramma antarctica] and crystal krill[Euphausia chrystallorophias]) responses to predation associated with the marginal ice zone (MIZ) of the McMurdo Sound, Antarctica, polynya. Prey abundance and habitat was sampled across a 30 × 15 km area by remotely-operated vehicle, and included locations that were accessible (ice edge) or inaccessible (solid fast ice) to air-breathing predators. Prey and habitat sampling coincided with bio-logging of Adélie penguins and observations of other air-breathing predators (penguins, seals, and whales), all of which were competing for the same prey. Adélie penguins dived deeper, and more frequently, near the ice edge. Lowered abundance of krill at the ice edge indicated they were depleted or were responding to increased predation and/or higher light levels along the ice edge. Penguin diet shifted increasingly to silverfish from krill during sampling, and was correlated with the arrival of krill-eating whales. Behaviorally-mediated, high trophic transfer characterizes the McMurdo Sound MIZ, and likely other MIZs, warranting more specific consideration in food web models and conservation efforts.

Understanding the effects of human exploitation on the genetic composition of wild populations is important for predicting species persistence and adaptive potential. We therefore investigated the genetic legacy of large-scale commercial harvesting by reconstructing, on a global scale, the recent demographic history of the Antarctic fur seal (Arctocephalus gazella), a species that was hunted to the brink of extinction by 18 th and 19 th century sealers. Molecular genetic data from over 2,000 individuals sampled from all eight major breeding locations across the species' circumpolar geographic distribution, show that at least four relict populations around Antarctica survived commercial hunting. Coalescent simulations suggest that all of these populations experienced severe bottlenecks down to effective population sizes of around 150-200. Nevertheless, comparably high levels of neutral genetic variability were retained as these declines are unlikely to have been strong enough to deplete allelic richness by more than around 15%. These findings suggest that even dramatic short-term declines need not necessarily result in major losses of diversity, and explain the apparent contradiction between the high genetic diversity of this species and its extreme exploitation history.

Competition for resources within a population can lead to niche partitioning between sexes, throughout ontogeny and among individuals, allowing con-specifics to co-exist. We aimed to quantify such partitioning in Antarctic fur seals, Arctocephalus gazella, breeding at South Georgia, which hosts ~95% of the world's population. Whiskers were collected from 20 adult males and 20 adult females and stable isotope ratios were quantified every 5 mm along the length of each whisker. Nitrogen isotope ratios (δ 15 N) were used as proxies for trophic position and carbon isotope ratios (δ 13 C) indicated foraging habitat. Sexual segregation was evident: δ 13 C values were significantly lower in males than females, indicating males spent more time foraging south of the Polar Front in maritime Antarctica. In males δ 13 C values declined with age, suggesting males spent more time foraging south throughout ontogeny. In females δ 13 C values revealed two main foraging strategies: 70% of females spent most time foraging south of the Polar Front and had similar δ 15 N values to males, while 30% of females spent most time foraging north of the Polar Front and had significantly higher δ 15 N values. This niche partitioning may relax competition and ultimately elevate population carrying capacity with implications for ecology, evolution and conservation.

Research using stable isotopes analysis (SIA) of carbon (δ 13 C) and nitrogen (δ 15 N) in blood components is lacking, because of the challenge of sample collection, processing, and storage in remote areas. There also is a paucity of information regarding the effect of tissue biochemical composition on isotopic ratios with few comparisons among taxa. We collected blood samples from shortfin mako sharks (n = 70; 2016) and Guadalupe fur seals (n = 25; 2017). All samples were centrifuged to obtain plasma from sharks and serum from the Guadalupe fur seals, and all the samples were prepared for SIA and analyzed using a Costech 4010 elemental analyzer interfaced with a Delta V Plus isotope ratio mass spectrometer. We found significant differences between plasma δ 13 C values of shortfin mako sharks (-17.6 ± 0.9‰) and serum of Guadalupe fur seals (-20.3 ± 1.2‰), but we did not find any differences for δ 15 N values between the two species. The differences in δ 13 C values between species are probably due to the specific blood composition and to the different biochemical characteristics and different adaptations within taxa. These findings highlight the importance of further research on the influence of biochemistry features on isotopic results, in this way a more accurate assessment will be possible for this factor, separating it from the dietary influences on stable isotopic values., (© 2019 Wiley Periodicals, Inc.)

In species exhibiting differential migration by sex and age, understanding what differences exist, and the adaptive reasons for these differences is critical for determining how demographic groups will respond to environmental variability and anthropogenic perturbations. We used satellite-telemetered movement and diving data to investigate differential migration and its ontogeny in a highly migratory North Pacific Ocean predator, the northern fur seal (Callorhinus ursinus; NFS), with a focus on understudied juvenile (1- to 2-year-old) animals. We instrumented 71 juvenile NFS in two years (2006-07 and 2007-08) at three major North American breeding sites and compared their migratory strategies with pups and adults. Although sexual dimorphism is strong in adult NFS, only weak differences in body mass between sexes were found in juveniles, which had similar body mass to pups (~3-4 months). However, unlike widely-dispersed pups, juvenile male and female NFS dispersed in different directions, and used different habitats characterized by distinct hydrography and prey assemblages during migration, similar to breeding adults. Juvenile diving behavior differed only modestly among habitats and between sexes, consistent with weak differences in body mass. Evidence of habitat sexual segregation by juvenile NFS contradicts previous hypotheses that physiological differences predominantly drive the ontogeny of differential migration.

Unihemispheric slow wave sleep (USWS) is a unique form of sleep in which one brain hemisphere maintains low voltage electrical activity indicative of waking while the opposite exhibits slow wave electrical activity indicative of sleep. USWS is present in several marine mammals and in some species of birds. One proposed biological function of USWS is to enable the animal to monitor the environment to detect predators or conspecifics. While asymmetrical eye state was often observed during behavioral sleep in birds and marine mammals, electrophysiological (electroencephalogram, EEG) correlates between the asymmetry of eye state and EEG of two cortical hemispheres have not been reliably established. This study examined the association between eye state and EEG activity during aquatic sleep in two subadult northern fur seals (Callorhinus ursinus), taking advantage of the simultaneous visibility of both eyes when the seals were in the prone position. We found that during USWS the eye contralateral to the sleeping hemisphere was closed on average 99.4±0.1% of the recording time. The eye contralateral to the waking hemisphere opened briefly for on average 1.9±0.1 sec with a rate of 8.2±1.0 per min. This eye was open on average 24.8±2.5% of the USWS time and it was closed no longer than 3 sec, on average 39.4±5.6% of the time. These data indicate that fur seals sleep in seawater by having intermittent visual monitoring. Our findings document the extent of visual monitoring of both eyes during USWS and support the idea that USWS allows intermittent visual vigilance. Thus, USWS serves two functions in the fur seal, facilitating movement and visual vigilance, which may also be the case in cetaceans., Competing Interests: The authors of this manuscript have read the journal's policy and have the following competing interests: LMM received support in the form of a salary from Utrish Dolphinarium-84 Ltd. OIL is affiliated with Utrish Dolphinarium-84 Ltd. but did not receive a salary conducting this study, data analysis and writing of the manuscript. There are no patents, products in development or marketed products to declare. This does not alter our adherence to PLOS ONE policies on sharing data and materials.

Here, we analyse changes throughout time in the isotopic niche of the Franciscana dolphin (Pontoporia blainvillei), the South American fur seal (Arctocephalus australis) and the South American sea lion (Otaria flavescens) from the Río de la Plata estuary and adjacent Atlantic Ocean to test the hypothesis that fishing may modify the diet of small-gape predators by reducing the average size of prey. The overall evidence, from stable isotope and stomach contents analyses, reveals major changes in resource partitioning between the three predators considered, mainly because of an increased access of Franciscana dolphins to juvenile demersal fishes. These results are consistent with the changes in the length distribution of demersal fish species resulting from fishing and suggest that Franciscana dolphin has been the most benefited species of the three marine mammal species considered because of its intermediate mouth gape. In conclusion, the impact of fishing on marine mammals goes beyond the simple reduction in prey biomass and is highly dependent on the mouth gape of the species involved.

Effective ecosystem-based management requires estimates of abundance and population trends of species of interest. Trend analyses are often limited due to sparse or short-term abundance estimates for populations that can be logistically difficult to monitor over time. Therefore it is critical to assess regularly the quality of the metrics in long-term monitoring programs. For a monitoring program to provide meaningful data and remain relevant, it needs to incorporate technological improvements and the changing requirements of stakeholders, while maintaining the integrity of the data. In this paper we critically examine the monitoring program for the Australian fur seal (AFS) Arctocephalus pusillus doriferus as an example of an ad-hoc monitoring program that was co-ordinated across multiple stakeholders as a range-wide census of live pups in the Austral summers of 2002, 2007 and 2013. This 5-yearly census, combined with historic counts at individual sites, successfully tracked increasing population trends as signs of population recovery up to 2007. The 2013 census identified the first reduction in AFS pup numbers (14,248 live pups, -4.2% change per annum since 2007), however we have limited information to understand this change. We analyse the trends at breeding colonies and perform a power analysis to critically examine the reliability of those trends. We then assess the gaps in the monitoring program and discuss how we may transition this surveillance style program to an adaptive monitoring program than can evolve over time and achieve its goals. The census results are used for ecosystem-based modelling for fisheries management and emergency response planning. The ultimate goal for this program is to obtain the data we need with minimal cost, effort and impact on the fur seals. In conclusion we identify the importance of power analyses for interpreting trends, the value of regularly assessing long-term monitoring programs and proper design so that adaptive monitoring principles can be applied., Competing Interests: The authors have declared that no competing interests exist.

The submersion of Late Pleistocene shorelines and poor organic preservation at many early archaeological sites obscure the earliest effects of humans on coastal resources in the Americas. We used collagen fingerprinting to identify bone fragments from middens at four California Channel Island sites that are among the oldest coastal sites in the Americas (~12,500-8,500 cal BP). We document Paleocoastal human predation of at least three marine mammal families/species, including northern elephant seals (Mirounga angustirostris), eared seals (Otariidae), and sea otters (Enhydra lutris). Otariids and elephant seals are abundant today along the Pacific Coast of North America, but elephant seals are rare in late Holocene (<1500 cal BP) archaeological sites. Our data support the hypotheses that: (1) marine mammals helped fuel the peopling of the Americas; (2) humans affected marine mammal biogeography millennia before the devastation caused by the historic fur and oil trade; and (3) the current abundance and distribution of recovering pinniped populations on the California Channel Islands may mirror a pre-human baseline.

Virtually all land mammals and birds have two sleep states: slow-wave sleep (SWS) and rapid eye movement (REM) sleep [1, 2]. After deprivation of REM sleep by repeated awakenings, mammals increase REM sleep time [3], supporting the idea that REM sleep is homeostatically regulated. Some evidence suggests that periods of REM sleep deprivation for a week or more cause physiological dysfunction and eventual death [4, 5]. However, separating the effects of REM sleep loss from the stress of repeated awakening is difficult [2, 6]. The northern fur seal (Callorhinus ursinus) is a semiaquatic mammal [7]. It can sleep on land and in seawater. The fur seal is unique in showing both the bilateral SWS seen in most mammals and the asymmetric sleep previously reported in cetaceans [8]. Here we show that when the fur seal stays in seawater, where it spends most of its life [7], it goes without or greatly reduces REM sleep for days or weeks. After this nearly complete elimination of REM, it displays minimal or no REM rebound upon returning to baseline conditions. Our data are consistent with the hypothesis that REM sleep may serve to reverse the reduced brain temperature and metabolism effects of bilateral nonREM sleep, a state that is greatly reduced when the fur seal is in the seawater, rather than REM sleep being directly homeostatically regulated. This can explain the absence of REM sleep in the dolphin and other cetaceans and its increasing proportion as the end of the sleep period approaches in humans and other mammals., (Published by Elsevier Ltd.)

Leopard seals are an important Antarctic apex predator that can affect marine ecosystems through local predation. Here we report on the successful use of micro geolocation logging sensor tags to track the movements, and activity, of four leopard seals for trips of between 142-446 days including one individual in two separate years. Whilst the sample size is small the results represent an advance in our limited knowledge of leopard seals. We show the longest periods of tracking of leopard seals' migratory behaviour between the pack ice, close to the Antarctic continent, and the sub-Antarctic island of South Georgia. It appears that these tracked animals migrate in a directed manner towards Bird Island and, during their residency, use this as a central place for foraging trips as well as exploiting the local penguin and seal populations. Movements to the South Orkney Islands were also recorded, similar to those observed in other predators in the region including the krill fishery. Analysis of habitat associations, taking into account location errors, indicated the tracked seals had an affinity for shallow shelf water and regions of sea ice. Wet and dry sensors revealed that seals hauled out for between 22 and 31% of the time with maximum of 74 hours and a median of between 9 and 11 hours. The longest period a seal remained in the water was between 13 and 25 days. Fitting GAMMs showed that haul out rates changed throughout the year with the highest values occurring during the summer which has implications for visual surveys. Peak haul out occurred around midday for the months between October and April but was more evenly spread across the day between May and September. The seals' movements between, and behaviour within, areas important to breeding populations of birds and other seals, coupled with the dynamics of the region's fisheries, shows an understanding of leopard seal ecology is vital in the management of the Southern Ocean resources., Competing Interests: The authors have declared that no competing interests exist.

Optimal foraging theory predicts that when food is plentiful all individuals should take a small range of preferred prey types, but as competition increases less preferred prey will be included in the diet. This dietary switching may not be uniform among individuals, which produces discrete dietary clusters. We tested this hypothesis for gentoo penguins at Bird Island, South Georgia, using stable isotope analysis and biologging. Competition, in the form of the density of foraging dives, increased markedly from incubation to chick-rearing owing to increased foraging effort. Birds responded behaviourally by exploiting a greater portion of the available foraging radius and increasing dive depths. Dietary niche width doubled and two discrete dietary clusters appeared; one comprising birds that consumed mostly krill and another that ate a greater proportion of demersal fish. There were no differences in morphology between the dietary classes, but birds in the fish class had a tendency to dive deeper, which suggests a behavioural basis for specialization. Our findings are consistent with the hypothesis that intra-specific competition expands the population's dietary niche width and drives divergence in diets among individuals.

Reproductive success is one of the central tenets of conservation management programs, yet the inability to study underlying physiological processes in a minimally-invasive manner and the unpredictable nature of wild animal populations leaves large gaps in our knowledge of factors critical to successful reproduction in wild species. This study integrated ultrasonography of the reproductive tract and analysis of reproductive hormones in 172 northern fur seals (Callorhinus ursinus) to identify intrinsic factors associated with reinitiating embryonic growth at the end of diapause. Within the first 3-4 weeks of active gestation, pregnant fur seals (n = 126) had a larger corpus luteum and fewer antral follicles than non-pregnant fur seals, or those still in diapause (n = 46). This suggests that the conceptus drives changes in ovarian status to convey its presence to the female. Morphological changes in the reproductive tract associated with pregnancy were not reflected in differences in endocrine profiles (estradiol, estrone, progesterone, and relaxin) between pregnant and non-pregnant individuals. Hormone concentrations correlated more strongly with calendar date than with the presence or size of the conceptus, demonstrating that none of these reproductive hormones were reliable markers for early pregnancy diagnosis. Instead, the northern fur seal's long diestrus may serve to reduce the probability of a temporal mismatch between corpus luteum regression and embryo implantation. Indeed, conception rates were high and confirmed rates of pregnancy loss were relatively low (11%). In this study, minimally-invasive ultrasonography was used in wild pinnipeds to detect very early pregnancy (embryonic vesicles >2 mm) in combination with ovarian and endocrine dynamics at the time of embryo implantation, shedding light on mechanisms for maternal recognition of pregnancy. This study is also the first to track whether these same animals carried the embryo to term, by observing fur seals during the birthing season the following year. Data do not support the notion that decreased pregnancy rates or higher pregnancy loss rates are major contributing factors to the northern fur seal's population decline., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Persistent Organic Pollutants (POPs) can cause toxic effects in many species which include endocrine dysfunction, immunotoxicity, developmental defects and neoplasia. Species dominating the upper trophic level are vulnerable to these effects due to bioaccumulation. In Bass Strait, the Australian fur seal (Arctocephalus pusillus doriferus) is an important top order predator and sentinel species for ecosystem health. An alopecia syndrome is seen at high prevalence in juvenile, female Australian fur seals at Lady Julia Percy Island, Victoria, Australia. Previous investigations suggest causality could be due to an endocrine-like toxicant. The alopecia syndrome has significance for thermoregulation and is a likely risk factor for mortality. Fur collected from case (alopecic) and control (unaffected) seals sampled at Lady Julia Percy Island were analysed for POPs. To investigate the utility of fur for monitoring POPs concentrations in pinnipeds, a comparison of POPs concentrations in the fur and blubber of Australian fur seals stranded along the Victorian coast was undertaken. The concentration of selected POPs including polychlorinated dibenzo-p-dioxins and dibenzofurans (PCDD/Fs), dioxin-like polychlorinated biphenyls (dl-PCBs), polybrominated diphenyl ethers (PBDEs) and perfluorooctane sulfonate/perfluorooctanoic acid (PFOS/PFOA) were determined in fur using either High Resolution Gas Chromatography-Mass Spectrometry or Liquid Chromatography-Mass Spectrometry. Results indicate detectable, and in some individuals, elevated levels of dl-PCBs, PCDD/Fs and PBDEs in juvenile fur seals sampled on Lady Julia Percy Island, with significantly higher levels of dl-PCBs in case compared to control seals. Elevated levels of dl-PCBs and PCDD/Fs were found in blubber samples collected from stranded fur seals with significant correlations between blubber and fur concentrations seen, particularly for dl-PCBs. This study discusses the significance of POPs concentrations in relation to the causality of an alopecia syndrome in the Australian fur seal, and assesses the utility of fur as a non-invasive biomarker to monitor POPs exposure in this sentinel species., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Direct measures of energy expenditure are difficult to obtain in marine mammals, and accelerometry may be a useful proxy. Recently its utility has been questioned as some analyses derived their measure of activity level by calculating the sum of accelerometry-based values and then comparing this summation to summed (total) energy expenditure (the so-called "time trap"). To test this hypothesis, we measured oxygen consumption of captive fur seals and sea lions wearing accelerometers during submerged swimming and calculated total and rate of energy expenditure. We compared these values with two potential proxies of energy expenditure derived from accelerometry data: flipper strokes and dynamic body acceleration (DBA). Total number of strokes, total DBA, and submergence time all predicted total oxygen consumption [Formula: see text] ml kg -1 ). However, both total DBA and total number of strokes were correlated with submergence time. Neither stroke rate nor mean DBA could predict the rate of oxygen consumption ([Formula: see text] ml min -1 kg -1 ). The relationship of total DBA and total strokes with total oxygen consumption is apparently a result of introducing a constant (time) into both sides of the relationship. This experimental evidence supports the conclusion that proxies derived from accelerometers cannot estimate the energy expenditure of marine mammals.

The pattern of sleep in the fur seal, a semiaquatic pinniped, has several striking behavioral and physiological adaptations that allow this species to inhabit both the land and water environment. These features include unihemispheric slow wave sleep (USWS, also being unihemispheric waking), the ability to maintain movement for stabilization of the sleep posture and to briefly open one eye while having a sleep electroencephalogram (EEG) in one hemisphere. In vivo microdialysis studies suggest that acetylcholine release is required for cortical activation during USWS, and that monoamines are not required for USWS. The need to breathe, to maintain efficient thermoregulation, and to avoid predation have shaped the sleep patterns in semiaquatic fur seals as in fully aquatic cetaceans., (Published by Elsevier Ltd.)

The efficiency with which individuals extract energy from their environment defines their survival and reproductive success, and thus their selective contribution to the population. Individuals that forage more efficiently (i.e., when energy gained exceeds energy expended) are likely to be more successful at raising viable offspring than individuals that forage less efficiently. Our goal was to test this prediction in large long-lived mammals under free-ranging conditions. To do so, we equipped 20 lactating Antarctic fur seals (Arctocephalus gazella) breeding on Kerguelen Island in the Southern Ocean with tags that recorded GPS locations, depth and tri-axial acceleration to determine at-sea behaviours and detailed time-activity budgets during their foraging trips. We also simultaneously measured energy spent at sea using the doubly-labeled water (DLW) method, and estimated the energy acquired while foraging from 1) type and energy content of prey species present in scat remains, and 2) numbers of prey capture attempts determined from head acceleration. Finally, we followed the growth of 36 pups from birth until weaning (of which 20 were the offspring of our 20 tracked mothers), and used the relative differences in body mass of pups at weaning as an index of first year survival and thus the reproductive success of their mothers. Our results show that females with greater foraging efficiencies produced relatively bigger pups at weaning. These mothers achieved greater foraging efficiency by extracting more energy per minute of diving rather than by reducing energy expenditure. This strategy also resulted in the females spending less time diving and less time overall at sea, which allowed them to deliver higher quality milk to their pups, or allowed their pups to suckle more frequently, or both. The linkage we demonstrate between reproductive success and the quality of individuals as foragers provides an individual-based quantitative framework to investigate how changes in the availability and accessibility of prey can affect fitness of animals.

Physiology may limit the ability for marine mammals to adapt to changing environments. Depth and duration of foraging dives are a function of total available oxygen stores, which theoretically increase as animals grow, and metabolic costs. To evaluate how physiology may influence the travelling costs for seals to foraging patches in the wild, we measured metabolic rates of a cross-section of New Zealand fur seals, Australian fur seals and Australian sea lions representing different foraging strategies, development stages, sexes and sizes. We report values for standard metabolic rate, active metabolic rate (obtained from submerged swimming), along with estimates of cost of transport (COT), measured via respirometry. We found a decline in mass-specific metabolic rate with increased duration of submerged swimming. For most seals mass-specific metabolic rate increased with speed and for all seals mass-specific COT decreased with speed. Mass-specific metabolic rate was higher for subadult than adult fur seals and sea lions, corresponding to an overall higher minimum COT. Some sex differences were also apparent, such that female Australian fur seals and Australian sea lions had higher mass-specific metabolic rates than males. There were no species differences in standard or active metabolic rates for adult males or females. The seals in our study appear to operate at their physiological optimum during submerged swimming. However, the higher metabolic rates of young and female fur seals and sea lions may limit their scope for increasing foraging effort during times of resource limitation.

In response to climate warming, a southward shift in productive frontal systems serving as the main foraging sites for many top predator species is likely to occur in Subantarctic areas. Central place foragers, such as seabirds and pinnipeds, are thus likely to cope with an increase in the distance between foraging locations and their land-based breeding colonies. Understanding how central place foragers should modify their foraging behavior in response to changes in prey accessibility appears crucial. A spatially explicit individual-based simulation model (Marine Central Place Forager Simulator (MarCPFS)), including bio-energetic components, was built to evaluate effects of possible changes in prey resources accessibility on individual performances and breeding success. The study was calibrated on a particular example: the Antarctic fur seal (Arctocephalus gazella), which alternates between oceanic areas in which females feed and the land-based colony in which they suckle their young over a 120 days rearing period. Our model shows the importance of the distance covered to feed and prey aggregation which appeared to be key factors to which animals are highly sensitive. Memorization and learning abilities also appear to be essential breeding success traits. Females were found to be most successful for intermediate levels of prey aggregation and short distance to the resource, resulting in optimal female body length. Increased distance to resources due to climate warming should hinder pups' growth and survival while female body length should increase.

Mitigating direct and indirect interactions between marine predators and fisheries is a motivating factor for ecosystem-based fisheries management (EBFM), especially where predators and fisheries compete for a shared resource. One difficulty in advancing EBFM is parameterizing clear functional responses of predators to indices of prey availability. Alternative characterizations of fishery-predator interactions may therefore benefit the implementation of EBFM. Telemetry data identify foraging areas used by predators and, therefore, represent critical information to mitigate potential competition between predators and fisheries. We analyzed six years (2009-2014) of telemetry data collected at Cape Shirreff, Livingston Island and Admiralty Bay, King George Island, Antarctica, on three species of Pygoscelid penguins and female Antarctic fur seals. In this region, all four species are primarily dependent on Antarctic krill. The tracking data demonstrate local movements near breeding colonies during the austral summer and dispersal from breeding colonies during the winter. We then assessed overlap between predators and the Antarctic krill fishery on a suite of spatiotemporal scales to examine how different data aggregations affect the extent and location of overlap. Concurrent overlap was observed on all spatiotemporal scales considered throughout the Antarctic Peninsula and South Orkney Islands region, including near tagging locations and in distant areas where recent fishing activity has concentrated. Overlap occurred at depths where mean krill densities were relatively high. Our results demonstrate that direct overlap of krill-dependent predators with the krill fishery on small spatiotemporal scales is relatively common throughout the Antarctic Peninsula region. As the krill fishery continues to develop and efforts to implement ecosystem-based management mature, indices of overlap may provide a useful metric for indicating where the risks of fishing are highest. A precautionary approach to allocating krill catches in space would be to avoid large increases in catch where overlap on small spatiotemporal scales is common., Competing Interests: The authors have declared that no competing interests exist.

Plastic debris in marine environments and its impact on wildlife species is becoming a problem of increasing concern. In pinnipeds, entanglements commonly consist of loops around the neck of non-biodegradable materials from fishing gear or commercial packaging, known as "neck collars". These entanglements can cause injuries, death by suffocation and starvation, and therefore they may add to the overall decrease in population. Our objective was to describe the entanglement of two species of otariids (Arctocephalus australis and Otaria flavescens) in the South West Atlantic Ocean. These two species have widely different population sizes and contrasting trends, being the O. flavescens population one order of magnitude lower in abundance with a negative population trend. A total number of 47 entangled individuals and the ingestion of a fishing sinker were recorded (A. australis: n = 26; O. flavescens: n = 22). For A. australis about 40% of the objects came from industrial fishing with which this species overlap their foraging areas, although also its lost or discarded gear can travel long distances. In O. flavescens 48% of observed injuries were very severe, which might indicate that they had been entangled for a long time. More than 60% of the objects came from artisanal and recreational fishing that operates within 5 nautical miles off the coast, which is probably related to coastal foraging habits of this species. Due to the frequent interaction between artisanal fisheries and O. flavescens, it is possible that entangled nets could be active gears. An important contribution to mitigate entanglements can be the development of education programs setting the scenario for effective communication, and exchange with involved fishermen to collect and recycle old fishing nets. Returning to natural fibers or replacement of the current materials used in fishing gear for biodegradable materials can also be a recommended mitigation measure., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Constructing activity budgets for marine animals when they are at sea and cannot be directly observed is challenging, but recent advances in bio-logging technology offer solutions to this problem. Accelerometers can potentially identify a wide range of behaviours for animals based on unique patterns of acceleration. However, when analysing data derived from accelerometers, there are many statistical techniques available which when applied to different data sets produce different classification accuracies. We investigated a selection of supervised machine learning methods for interpreting behavioural data from captive otariids (fur seals and sea lions). We conducted controlled experiments with 12 seals, where their behaviours were filmed while they were wearing 3-axis accelerometers. From video we identified 26 behaviours that could be grouped into one of four categories (foraging, resting, travelling and grooming) representing key behaviour states for wild seals. We used data from 10 seals to train four predictive classification models: stochastic gradient boosting (GBM), random forests, support vector machine using four different kernels and a baseline model: penalised logistic regression. We then took the best parameters from each model and cross-validated the results on the two seals unseen so far. We also investigated the influence of feature statistics (describing some characteristic of the seal), testing the models both with and without these. Cross-validation accuracies were lower than training accuracy, but the SVM with a polynomial kernel was still able to classify seal behaviour with high accuracy (>70%). Adding feature statistics improved accuracies across all models tested. Most categories of behaviour -resting, grooming and feeding-were all predicted with reasonable accuracy (52-81%) by the SVM while travelling was poorly categorised (31-41%). These results show that model selection is important when classifying behaviour and that by using animal characteristics we can strengthen the overall accuracy., Competing Interests: The authors have declared that no competing interests exist.

Investigating the ontogeny of niche differentiation enables to determine at which life-stages sexual segregation arises, providing insights into the main factors driving resource partitioning. We investigated the ontogeny of foraging ecology in Antarctic fur seals (Arctocephalus gazella), a highly dimorphic species with contrasting breeding strategies between sexes. Sequential δ(13)C and δ(15)N values of whiskers provided a longitudinal proxy of the foraging niche throughout the whole life of seals, from weaning, when size dimorphism is minimal to the age of 5. Females exhibited an early-life ontogenetic shift, from a total segregation during their first year at-sea, to a similar isotopic niche as breeding females as early as age 2. In contrast, males showed a progressive change in isotopic niche throughout their development such that 5-year-old males did not share the same niche as territorial bulls. Interestingly, males and females segregated straight after weaning with males appearing to feed in more southerly habitats than females. This spatial segregation was of similar amplitude as observed in breeding adults and was maintained throughout development. Such early-life niche differentiation is an unusual pattern and indicates size dimorphism and breeding constraints do not directly drive sexual segregation contrary to what has been assumed in otariid seals.

Marine mammals have lost the ability to express S-cone opsin, and possess only one type of M/L-cone in addition to numerous rods. As they are cone monochromats they should be color blind. However, early behavioral experiments with fur seals and sea lions indicated discrimination ability between many shades of grey and blue or green. On the other hand, most recent training experiments with harbor seals under "mesopic" conditions demonstrated rod based color blindness (Scholtyssek et al., 2015). In our experiments we trained two harbor seals (Phoca vitulina) and two South African fur seals (Arctocephalus pusillus) with surface colors under photopic conditions. The seals had to detect a triangle on grey background shown on one of three test fields while the other two test fields were homogeneously grey. In a first series of experiments we determined brightness detection. We found a luminance contrast of >3% sufficient for correctly choosing the triangle. In the tests for color vision the triangle was blue, green or yellow in grey surround. The results show that the animals could see the colored triangle despite minimal or zero brightness contrast. Thus, seals have color vision based on the contribution of cones and rods even in bright daylight., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

The abundance of California sea lions (Zalophus californianus) (CSLs) and Guadalupe fur seals (Arctocephalus philippii townsendi) (GFSs) from the San Benito Archipelago (SBA) was determined through nine monthly surveys in 2014-2015. Assessment of their foraging habits was examined based on the isotopic analysis of pups (maternal indicators) (SIAR/SIBER-R). Environmental variability between 2014 and 2015 was also analyzed, in terms of sea surface temperature (SST) and chlorophyll (Chl-a) concentration. Both otariids reached their highest abundance in July of both years; however, relative to 2014, the 2015 survey showed a 59.7% decline in the total GFS abundance and a 42.9% decrease of GFS pups, while total CSL abundance decreased 52.0% and CSL pup presence decreased in 61.7%. All monthly surveys for both otariids showed a similar trend (>50% decrease in 2015). Compared to 2014, the 2015 GFSs isotopic niche was three times larger (2.0 in 2015, 0.6 in 2014) and the δ13C was significantly lower. CSLs also showed significantly lower δ13C and higher δ15N in 2015. Interannual segregation was greater for CSLs, and their pup body mass was also significantly lower during the 2015 breeding season (mean = 8.7 kg) than in the same season of 2014 (mean = 9.9 kg). The decrease in δ13C for both otariids reflected a more oceanic foraging; most likely associated with the decline in primary productivity in surrounding areas to the SBA, related to a higher SST caused by the 2015 ENSO, with a subsequent increase in foraging effort. These would explain the fewer observed individuals on land, especially pups, which showed diminished body condition (CSLs). This study highlights the importance of marine mammals as sentinel species that respond dynamically to changes in environment, providing valuable information on the effect of ENSO on pinnipeds in Mexican waters.

Foraging behaviour of marine top predators is increasingly being used to identify areas of ecological importance. This is largely enabled by the ability of many such species to forage extensively in search of prey that is often concentrated in oceanographically productive areas. To identify important habitat in the Southern Indian Ocean within and around South Africa's Prince Edward Islands' Marine Protected Area (MPA), satellite transmitters were deployed on 12 lactating Subantarctic fur seals Arctocephalus tropicalis at Prince Edward Island (PEI) itself. Switching state space models were employed to correct ARGOS tracks and estimate behavioural states for locations along predicted tracks, namely travelling or area restricted search (ARS). A random forest model showed that distance from the study colony, longitude and distance from the Subantarctic Front were the most important predictors of suitable foraging habitat (inferred from ARS). Model-predicted suitable habitat occurred within the MPA in relatively close access to the colony during summer and autumn, but shifted northwards concurrently with frontal movements in winter and spring. The association of ARS with the MPA during summer-autumn was highly significant, highlighting the effectiveness of the recently declared reserve's design for capturing suitable foraging habitat for this and probably other marine top predator species.

Study Objectives: Our understanding of the role of neurotransmitters in the control of the electroencephalogram (EEG) has been entirely based on studies of animals with bilateral sleep. The study of animals with unihemispheric sleep presents the opportunity of separating the neurochemical substrates of waking and sleep EEG from the systemic, bilateral correlates of sleep and waking states., Methods: The release of histamine (HI), norepinephrine (NE), and serotonin (5HT) in cortical and subcortical areas (hypothalamus, thalamus and caudate nucleus) was measured in unrestrained northern fur seals (Callorhinus ursinus) using in vivo microdialysis, in combination with, polygraphic recording of EEG, electrooculogram, and neck electromyogram., Results: The pattern of cortical and subcortical HI, NE, and 5HT release in fur seals is similar during bilaterally symmetrical states: highest in active waking, reduced in quiet waking and bilateral slow wave sleep, and lowest in rapid eye movement (REM) sleep. Cortical and subcortical HI, NE, and 5HT release in seals is highly elevated during certain waking stimuli and behaviors, such as being sprayed with water and feeding. However, in contrast to acetylcholine (ACh), which we have previously studied, the release of HI, NE, and 5HT during unihemispheric sleep is not lateralized in the fur seal., Conclusions: Among the studied neurotransmitters most strongly implicated in waking control, only ACh release is asymmetric in unihemispheric sleep and waking, being greatly increased on the activated side of the brain., Commentary: A commentary on this article appears in this issue on page 491., (© 2016 Associated Professional Sleep Societies, LLC.)

Estimating the degree of individual specialisation is likely to be sensitive to the methods used, as they record individuals' resource use over different time-periods. We combined animal-borne video cameras, GPS/TDR loggers and stable isotope values of plasma, red cells and sub-sampled whiskers to investigate individual foraging specialisation in female Australian fur seals (Arctocephalus pusillus doriferus) over various timescales. Combining these methods enabled us to (1) provide quantitative information on individuals' diet, allowing the identification of prey, (2) infer the temporal consistency of individual specialisation, and (3) assess how different methods and timescales affect our estimation of the degree of specialisation. Short-term inter-individual variation in diet was observed in the video data (mean pairwise overlap = 0.60), with the sampled population being composed of both generalist and specialist individuals (nested network). However, the brevity of the temporal window is likely to artificially increase the level of specialisation by not recording the entire diet of seals. Indeed, the correlation in isotopic values was tighter between the red cells and whiskers (mid- to long-term foraging ecology) than between plasma and red cells (short- to mid-term) (R(2) = 0.93-0.73 vs. 0.55-0.41). δ(13)C and δ(15)N values of whiskers confirmed the temporal consistency of individual specialisation. Variation in isotopic niche was consistent across seasons and years, indicating long-term habitat (WIC/TNW = 0.28) and dietary (WIC/TNW = 0.39) specialisation. The results also highlight time-averaging issues (under-estimation of the degree of specialisation) when calculating individual specialisation indices over long time-periods, so that no single timescale may provide a complete and accurate picture, emphasising the benefits of using complementary methods.

A species, according to the biological concept, is a natural group of potentially interbreeding individuals isolated by diverse mechanisms. Hybridization is considered the production of offspring resulting from the interbreeding of two genetically distinct taxa. It has been documented in over 10% of wild animals, and at least in 34 cases for Artic marine mammals. In Otariids, intergeneric hybridization has been reported though neither confirming it through genetic analyses nor presenting evidence of fertile offspring. In this study, we report the finding of a hybrid adult female between a South American fur seal (Arctocephalus australis) and a South American sea lion (Otaria byronia), and its offspring, a male pup, in Uruguay. Further based on morphological constraints and breeding seasons, sex-biased hybridization between the two species is hypothesized. Morphological and genetic (nuclear and mitochondrial) results confirm de hybrid nature of the female-pup pair. Here we discuss a genetic dilution effect, considering other hybridization events must be occurring, and how isolation mechanisms could be circumvented. Moreover, the results obtained from stable isotope analysis suggest feeding habits may be a trait transmitted maternally, leading to consider broader issues regarding hybridization as an evolutionary innovation phenomenon., (© 2016 Wiley Periodicals, Inc.)

Understanding temporal patterns of marine mammal occurrence is useful for establishing conservation strategies. We used a 38 yr-long dataset spanning 1976 to 2013 to describe temporal patterns and trends in marine mammal strandings along a subtropical stretch of the east coast of South America. This region is influenced by a transitional zone between tropical and temperate waters and is considered an important fishing ground off Brazil. Generalized Additive Models were used to evaluate the temporal stranding patterns of the most frequently stranded species. Forty species were documented in 12,540 stranding events. Franciscana (n = 4,574), South American fur seal, (n = 3,419), South American sea lion (n = 2,049), bottlenose dolphins (n = 293) and subantarctic fur seal (n = 219) were the most frequently stranded marine mammals. The seasonality of strandings of franciscana and bottlenose dolphin coincided with periods of higher fishing effort and strandings of South American and subantarctic fur seals with post-reproductive dispersal. For South American sea lion the seasonality of strandings is associated with both fishing effort and post-reproductive dispersal. Some clear seasonal patterns were associated with occurrence of cold- (e.g. subantarctic fur seal) and warm-water (e.g. rough-toothed dolphin) species in winter and summer, respectively. Inter-annual increases in stranding rate were observed for franciscana and South American fur seal and these are likely related to increased fishing effort and population growth, respectively. For subantarctic fur seal the stranding rate showed a slight decline while for bottlenose dolphin it remained steady. No significant year to year variation in stranding rate was observed for South American sea lion. The slight decrease in frequency of temperate/polar marine mammals and the increased occurrence of subtropical/tropical species since the late 1990s might be associated with environmental changes linked to climate change. This long-term study indicates that temporal stranding patterns of marine mammals might be explained by either fishing-related or environmental factors.

Feeding performance studies can address questions relevant to feeding ecology and evolution. Our current understanding of feeding mechanisms for aquatic mammals is poor. Therefore, we characterized the feeding kinematics and performance of five Steller sea lions (Eumetopias jubatus) and six northern fur seals (Callorhinus ursinus). We tested the hypotheses that both species use suction as their primary feeding mode, and that rapid jaw opening was related to suction generation. Steller sea lions used suction as their primary feeding mode, but also used a biting feeding mode. In contrast, northern fur seals only used a biting feeding mode. Kinematic profiles of Steller sea lions were all indicative of suction feeding (i.e. a small gape, small gape angle, large depression of the hyolingual apparatus and lip pursing). However, jaw opening as measured by gape angle opening velocity (GAOV) was relatively slow in Steller sea lions. In contrast to Steller sea lions, the GAOV of northern fur seals was extremely fast, but their kinematic profiles indicated a biting feeding mode (i.e. northern fur seals exhibited a greater gape, a greater gape angle and minimal depression of the hyolingual apparatus compared with Steller sea lions). Steller sea lions produced both subambient and suprambient pressures at 45 kPa. In contrast, northern fur seals produced no detectable pressure measurements. Steller sea lions have a broader feeding repertoire than northern fur seals, which likely enables them to feed on a greater variety of prey, in more diverse habitats. Based on the basal phylogenetic position of northern fur seals, craniodental morphological data of the Callorhinus lineage, and the performance data provided in this study, we suggest that northern fur seals may be exhibiting their ancestral feeding mode., (© 2015. Published by The Company of Biologists Ltd.)

Rationale: In mammals including humans, mother-to-offspring transfer of nutrients has been the focus of several isotopic studies. Measurement of δ(13) C and δ(15) N values were mainly conducted on easily sampled tissues such as blood and hair that allow the calculation of apparent discrimination factors (∆(13) C and ∆(15) N) between offspring and maternal tissues. Quantifying real ∆(13) C and ∆(15) N values requires the measurement of the δ(13) C and δ(15) N values of milk, the exclusive food of newborns. Surprisingly, little isotopic information is available on milk and its biochemical components (lipids and proteins)., Methods: Paired blood and milk samples from 10 lactating females and their pups were collected from two otariid species, the Antarctic and subantarctic fur seals. Tissue δ(13) C and δ(15) N values were measured using continuous-flow isotope ratio mass spectrometry (CFIRMS) on maternal and offspring blood, and on whole milk, lipid-free milk and milk lipids, thus allowing the calculation and comparison of apparent (maternal blood to offspring blood) and real (lipid-free milk to offspring blood) ∆(13) C and ∆(15) N values., Results: In both fur seal species, the apparent ∆(13) C values averaged ~0.0 ‰. Lipid-free milk was slightly (13) C-depleted compared with both maternal and pup blood and it was strongly (13) C-enriched (~6.3 ‰) compared with milk lipids. In contrast, the apparent and real ∆(15) N values averaged 1.2-1.4 and 2.6-3.0 ‰, respectively, the differences being explained by the ~1.5 ‰ lower milk δ(15) N values than those of maternal blood., Conclusions: In fur seals, the low apparent ∆(15) N translated into a higher real ∆(15) N value, amounting to a full trophic level, which is in agreement with the almost never verified hypothesis that (15) N differences between mothers and their offsprings should reflect one complete trophic level. The study highlights the need to measure milk isotopic values to disentangle the nutritional mother-to-offspring relationships., (Copyright © 2015 John Wiley & Sons, Ltd.)

While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ13C values. However, males exhibited greater δ13C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ15N values. Yet, males displayed higher δ15N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ15N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ13C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.

The foraging strategy of many animals is thought to be determined by their past experiences. However, few empirical studies have investigated whether this is true in diving animals. We recorded three-dimensional movements and mouth-opening events from three Antarctic fur seals during their foraging trips to examine how they adapt their behaviour based on past experience--continuing to search for prey in the same area or moving to search in a different place. Each dive cycle was divided into a transit phase and a feeding phase. The linear horizontal distance travelled after feeding phases in each dive was affected by the mouth-opening rate during the previous 244 s, which typically covered two to three dive cycles. The linear distance travelled tended to be shorter when the mouth-opening rate in the previous 244 s was higher, i.e. seals tended to stay in the same areas with high prey-encounter rates. These results indicate that Antarctic fur seals follow decision-making strategies based on the past foraging experience over time periods longer than the immediately preceding dive., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)